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SABLES: GENETICS AND MYTHS
by Fred Lanting
Copyright October, 2008 -
Some time ago, a GSD breeder-judge in Germany wrote an article that was carried on the Internet, entitled "The Sable Shepherd… A Museum Piece?" (liberal translation). Several people have asked me about some of the statements in it, and I thought a magazine or website piece might be the easiest way to answer.
The color pattern we in America call "sable" is called "grau" (gray) in Germany. Neither word is fully accurate, but experienced dog people know what is meant. The strict translation of what is possibly originally Latin, then Russian-Scandinavian in origin, but is found in variations in many languages including "sable" in English and French, is "black". Actually, even before that meaning, it was the name of a small glossy-black weasel or ermine found in far-northern Europe and Asia. From the Latin zobola, Russian soboli, Scandinavian sabel, and German zobel, we get our word, but not our meaning. It brings a different image in other breeds, such as the Collie, Basenji, and Sheltie where it refers to a reddish-yellow dog. Even in the Pembroke Welsh Corgi, the sable dog gives mostly an orange impression. In the Shar-Pei, we see a color that looks like the GSD sable, but has a different genetic constitution. There is even the white GSD, some of which appear to be genetically modified sables, but certainly without any black.
Let’s define a couple more terms before proceeding. Everyone knows what is usually meant by the word "color": the reflection or refraction of certain wavelengths of light that tell us the apple is green, the traffic light is red, or the rainbow has seven more or less different hues. But "color" is often the term used when GSD people talk about black-and-tans, blacks, bi-colored dogs, or sables. For that meaning we really should use the term "color pattern", because some of these dogs may have more than one color or hue in the coat: shades of red/yellow/brown, black, white, and "dilutes" (blue and liver). The German word for what we call "tan" is "gelbe" (yellow). While "braun" (what Americans often braggingly like to call "red") is also seen in the Ahnentafeln (SV registration-pedigree), it is genetically and chemically the same as yellow or red, altered with hue and intensity modifier genes.
A hair bulb can manufacture more than one type of melanin, and can alternate production in such a way that some hairs, such as most of those on a sable dog, are dark tipped, followed by a lighter midpiece and an even lighter base (or perhaps a dark base). There may be two shades of yellow: one reddish and the other cream, on one shaft. Sometimes the phaeomelanin (this form produces the non-dark pigment) is concentrated in the tip, and the eumelanin (which produces the black, blue, or liver) is in the base, though not often. The banding or alternating between dark and light sections results in a beautiful variety of colorations in the breed, especially around the neck, withers, and shoulders. The definition of a sable pattern should refer to the guard hairs being tipped with black. The more hairs and the longer that tipping, the darker the dog. The richer the phaeomelanin-influenced parts (undercoat and lower/ventral/leg portions), the more the dog is called a "red sable" instead of a "gray sable" (or faded). The "black sables" are those with much tipping but not always much red underneath.
So-called "golden sable" resulting from good brown lower parts of the guard hairs but not as much black tipping as some other dogs pictured in this article.
Agouti is a term borrowed from genetics studies on that type of rat. It refers to the banded‑hair coloration or pattern seen in Elkhounds, wolves, and sable German Shepherds, but can also be seen in varying amounts in the neck, shoulder, tail, croup and border markings in saddle‑marked dogs. Some saddle-marked B&T GSDs have so much grizzling in the black blanket or so much undercoat showing, that they are hard to distinguish from sables, and some sables that carry the partially-hidden B&T-saddle pattern recessive have so much of that recessive showing through that they look like washed-out B&Ts. At one chromosome’s locus A, the German Shepherd Dog has a pair of genes, one on each of the chromosome pair, which together determine the major markings we call "pattern". This pair may have two of the same or two different alleles, one more dominant and the other recessive to it.
Ay, sable — The superscript y is used in genetic parlance to refer to the yellow pigment that can dominate the appearance of the tan‑sable dog, even though a true sable should have those hairs tipped with black. That black appearance has been lost in many breeds through selective breeding. In other breeds, where a more dominant allele for self-color (solid) exists, the sable or yellow pattern is ay (lower-case a, because the capital letter refers to the most dominant allele in the series). For example, the black Lab is As while the yellow Lab is ay. The color lying under the black tips on a GSD’s coat, for example, is due to other genes. A sable is generally held to mean a dog with black-tipped guard hairs, this banding being obvious over most of the dog’s surface, especially the dorsal parts. In those other breeds I mentioned, the word is used variously to refer to a yellow dog regardless of the presence of any black tipping. Many or most "sable" Collies and Corgis have lost the genes calling for expression of black in the hair. There is also a "white sable", but that subject is treated in a different article.
as, saddle — The saddle pattern gene is below Ay in order of dominance. The use of lower case shows it is not as strong in its influence over pattern as the sable gene, but we know it is dominant over other alleles in the series. The saddle pattern has been the most popular in the past 50 years. Whether, as the Germans describe it, the base color is black and the tan portions are the markings, or whether (as some of us think of it) the base color is brown/tan and the saddle, muzzle, and head markings are the areas of added pigmentation, it's moot, like arguing about the color of a zebra.
at , bicolor — Again, this pattern, resembling the coloration of a B&T Coonhound or Doberman Pinscher, is best discussed in a separate article. Some think it belongs on a different chromosome locus.
Incomplete dominance — In the A series, incomplete dominance of one A series allele over a lower one is often, though not always, visible to the experienced eye. A heterozygous Ay as sable (i.e., the genes in the pattern pair not identical, and we would say he carries a recessive for saddle pattern) will have a shadow of a saddle with slightly more dark‑tipped hairs (or longer black band on those hairs) than the homozygous Ay Ay sable has in the areas commonly marked black in the saddle dog. If the sable has a hypostatic (recessive) gene (at the E locus?) for solid black, such a dog is usually considerably darker in overall appearance than are his lighter gray or golden sable relatives. Karilea's Cito, Dipadon's Dasher, and other sables that carried the black factor and were capable of producing solid blacks had this dark look. Many "working-lines" German Shepherd Dogs carry the black or bi-color recessive (or both?) and have a very dark appearance. It may well be that most extensively-covered very dark sables carry the bicolor recessive, and/or the black factor at the E locus as well. This may be especially true in the working lines, where these three patterns have not been pushed out of the picture by the saddle-marked dogs as has happened in the breed show ring. For greater detail on genetics of coat color and other factors, get a copy of Fred Lanting's Total GSD
Since beginners are often confused as to what will happen if they breed a certain bitch to a certain dog, let’s put some dogs together on paper and see what you get. As an example, the homozygous sable, with both A series alleles (genes in the pair) being Ay, is bred to the homozygous B&T saddle, as as. Regardless of other factors such as depth or distribution of pigment, the Ay Ay X as as (sable-saddle cross) will give a litter of 100 percent sable phenotype, though possibly not as distinctly patterned as the one parent's, and all the pups will carry the as as a recessive. Very few homozygous sables seem to be running around, so most of the ones you see will produce some B&T pups, even when bred to another heterozygous sable. The variations in shades and markings are influenced by "modifier genes" such as E, Em, e, or eb which govern the extension of black pigment to a mask, and influence on what percentage of the body that pigment will be used. E alleles appear to act more as "advisors" to the A series. There was a nice article on masks and patterns, by the "sable" (blonde, not black-tipped) Linda Shaw, in the July/August 2000 Schutzhund-USA issue.
The Famed Scotland-born Dunmonaidh Junker
Sable Prima Zorba’s B&T recessive shows through
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Fred Lanting is an internationally respected show judge, approved by many registries as an all-breed judge, has judged numerous countries’ Sieger Shows and Landesgruppen events, and has many years experience with SV. He presents seminars and consults worldwide on such topics as Gait-&-Structure, HD and Other Orthopedic Disorders, Anatomy, Training Techniques, and The GSD. He conducts annual non-profit sightseeing tours of Europe, centered on the Sieger Show (biggest breed show in the world) and BSP.
All Things Canine -- consulting division, Willow Wood Services Phone: 256-498-3319 Fax: 256-498-3311 E-mail mr.gsd@netscape.com
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